Lepidoblepharis emberawoundule, Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, Köhler, Gunther, Carrizo, Arcadio & Lotzkat, Sebastian, 2015

Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, Köhler, Gunther, Carrizo, Arcadio & Lotzkat, Sebastian, 2015, Revision of the genus Lepidoblepharis (Reptilia: Squamata: Sphaerodactylidae) in Central America, with the description of three new species, Zootaxa 3994 (2), pp. 187-221 : 194-196

Lepidoblepharis emberawoundule sp. nov.

Figs. 3–7 , 10 .

Lepidoblepharis xanthostigma : all in part. (referring to certain populations in eastern Panama): Auth (1994); Young et al. (1999); Ibáñez et al. (2001); Köhler (2001: Fig. 172; 2008: Fig. 136); Jaramillo et al. (2010).

Holotype. Adult male SMF 50968, original field number AB 963 ( Fig. 10 ), collected from leaf-litter at La Cascada trail, Burbayar private reserve (9.31837°N, 79.00266°W, 360 m elev.), Cartí, Narganá, Comarca Guna Yala, Panama, on 26 November 2013 at 23:40 hrs, by Abel Batista and Konrad Mebert.

Paratypes. Three adult males, two adult females, and one juvenile, all from Panama. Three males ( SMF 81950–52) from Nusagandí field station and two females ( SMF 81953–54) from the nearby Sendero Nusagandí, Comarca Guna Yala, collected 14–17 April 2000; one juvenile ( MHCH 2952) from Río Terable, El Llano, Chepo, Panama, collected on 27 November 2012; see Appendix I for locality details.

Referred specimens. MHCH 2951, 2957; SMF 50969–70; FMNH 170029, 170042–45; see Appendix I for locality details.

Diagnosis. Lepidoblepharis emberawoundule (our sp. nov. 1) is characterized by the following combination of characters: (1) dorsal body scales small, granular, and juxtaposed, ventral scales large, cycloid, flat, and imbricate; (2) scales on head small and granular; (3) 3–4 (3.1 ± 0.3) postrostral scales; (4) a vaguely M-shaped posterior mental border with two paramedian clefts; (5) 3–7 (5.1 ± 1.04) postmentals, larger than the posteriorly adjacent scales on chin; (6) lamellae under fourth toe 6–9 (7.9 ± 0.85), lamellae under fourth finger 5–8 (6.7 ± 1.01); (7) median subcaudals conspicuously wider than long, but their width less than twice the width of the laterally adjacent scales or their own length, with straight or rounded posterior margins, arranged in a regular tail sequence of 1'1''; (8) ventral escutcheon consisting of 38–61 (48.4 ± 8.85) scales, 6–7 (6.8 ± 0.45) scales long and 10–13 (11.6 ± 1.14) wide; (9) subfemoral escutcheon consisting of 4–5 (4.4 ± 0.52) well-discernible scales per thigh arranged in a single row ( Fig. 5 A); (10) 16–20 (18.2 ± 1.17) longitudinal rows of ventral scales at midbody; (11) 52–74 (61.8 ± 7.61) longitudinal rows of dorsal scales at midbody; (12) bilobate hemipenis, with a third lobule rising from the pedicel ( Fig. 6 A–B); (13) SVL 21–30 (25 ± 2.07) mm.

Comparison with other species of the genus. Lepidoblepharis emberawoundule can be differentiated from many species in the genus by its small size and its low number of lamellae under the fourth toe and finger ( Figs. 3– 4 ). In the following, we provide comparisons to all other species within the genus, with the characteristics for L. emberawoundule in parentheses. Lepidoblepharis emberawoundule can be distinguished from the Panamanian species L. xanthostigma , L. sanctaemartae , Lepidoblepharis sp. nov. 2 (described below), and Lepidoblepharis sp. nov. 3 (described below) by uncorrected genetic p-distance (10–26% in 16S mtDNA between individuals).

Lepidoblepharis xanthostigma has greatly enlarged median subcaudal scales (slightly enlarged), and 12–16 lamellae under its fourth toe (6–9). Lepidoblepharis sanctaemartae has large, flat, imbricate dorsal body scales (small granular scales). Lepidoblepharis sp. nov. 2 (described below) has 13 lamellae under its fourth toe (6–9) and 11 under its fourth finger (5–8). Lepidoblepharis sp. nov. 3 (described below) has 4–5 lamellae under its fourth finger (5–8) and a unique lamellar configuration with 1–3 proximal lamellae per digit greatly enlarged, i.e., about 3–4 times longer than any of the remaining lamellae, the ventral escutcheon consisting of 61–68 scales (38–61), and no discernible subfemoral escutcheon (4–5 discernible subfemoral escutcheon scales per thigh). To date, seven species of the genus Lepidoblepharis have been reported to possess ten or fewer lamellae under the fourth toe, i.e., to be short-toed. Two of these, L. miyatai Lamar 1985 and L. sanctaemartae , possess large, flat, and imbricate dorsal scales (small, granular, and juxtaposed dorsals). Three others, Lepidoblepharis buchwaldi Werner 1910 , L. montecanoensis Markezich & Taphorn 1994 , and L. williamsi Ayala & Serna 1986 , can be readily distinguished from L. emberawoundule because the inspection of the illustrations and photographs available for the respective holotypes showed clear differences between the species: In L. buchwaldi , the enlarged subcaudals are much wider than long and at least twice as wide as the laterally adjacent subcaudals (less than twice as wide as they are long or as the neighboring subcaudals are wide), the dorsal tail scales are small, i.e., less than twice the size of the dorsal body scales (twice or more the size of the dorsal body scales), and the posterior border of the mental has a single median cleft (two paramedian clefts). Lepidoblepharis montecanoensis is a very small species with a SVL of 18–21 mm (21–30), and lacks defined occipital marks in males (two well defined occipital marks in males) as well as distinctly enlarged median subcaudals (median subcaudals distinctly enlarged). Lepidoblepharis williamsi also lacks enlarged median subcaudal scales (median subcaudals distinctly enlarged), and has only 25–40 ventral escutcheon scales (38–61). The holotype of L. peraccae Boulenger 1908 has eight lamellae under the fourth finger (5–8) and ten under the fourth toe (6–9), its plantar and palmar scales have ovoid and strongly imbricate posterior borders (those scales small, rounded, and juxtaposed; Fig. 7 ). The holotype of L. microlepis ( Noble 1923) is very similar to L. emberawoundule , but differs in the scalation of the chin region and the ventral tail surface ( Fig. 7 ). The posterior margin of its mental is V-shaped and lacks conspicuous clefts (posterior margin M-shaped, i.e., slightly convex in the middle, with two conspicuous paramedian clefts), there are six postmentals, with one medial postmental greatly enlarged and two neighboring scales slightly enlarged (3–7 postmentals, median scales slightly larger than the others), and the posteriorly adjacent chin scales are small and conical (small and flat, some slightly pointed, and juxtaposed, Figs. 3 , 7 ). Most decisively, each of the slightly enlarged subcaudal scales of the holotype of L. microlepis is bordered laterodistally by only one scale, leading to a regular tail sequence of 1'1' (the larger of the enlarged subcaudals bordered laterodistally by two scales, the smaller ones by one, forming a regular tail sequence of 1'1''; see Fig. 7 E–F). The remaining species of the genus, i.e., L colombianus Mechler 1968 , L conolepis Avila-Pires 2001 , L. duolepis Ayala & Castro 1983 , L. festae Peracca 1897 , L. grandis Miyata 1985 , L. heyerorum Vanzolini 1978 , L. hoogmoedi Avila-Pires 1995 , L. intermedius Boulenger 1914 , and L. ruthveni Parker 1926 are long-toed with eleven or more lamellae under the fourth toe (6–9 in L. emberawoundule ).

Description of the holotype. Variation among the entire type series is given in parentheses for selected characters (see Table 1 for details and variation among all examined specimens). Adult male as judged by everted hemipenes; SVL 26 mm (21–27 mm), TL 27 mm (measured while the now broken tail was still intact) (18–27 mm), HL 5.6 mm (5.0– 5.6 mm), HW 3.8 mm (3.2–4.4 mm), forelimbs 5.2 mm, hind limbs 8.6 mm, shank 3.7 mm; rostral large, clearly visible from above, with a shallow, horseshoe-shaped posterior depression and a long median cleft; postrostrals including supranasals three (3–4), the median ones smaller than supranasals, and indenting the rostral, the median postrostral about the same size as the posterior scales on snout; postnasals two (1–2), both about the same size as posteriorly adjacent loreal scales; scales on snout small and smooth, 20 (18–21) scales across snout between anterior sutures of second SPLs; loreal scales juxtaposed, elevated, and rounded, 7 (6–7) on a longitudinal line between postnasals and orbit; scales on top of head small, granular, juxtaposed, generally pointing upward, about as half as large as those on the middle area of snout; superciliary flap with two enlarged scales on anterior border, of which the first is slightly longer than the second, followed by four small scales; supralabials three (3–4), posteriormost one below center of eye; ear-opening small, oval, in oblique orientation; mental large, posterior margin slightly convex in the middle, with two small clefts bordering this convexity, resulting M-shaped; postmentals 7 (4–7), median scales slightly larger than the others, postmentals larger than the posteriorly adjacent chin scales ( Figs. 3 , 7 ); scales on chin small and juxtaposed, most of them flat but some slightly pointed, on posterior region granular, approximately vertical in position, with a slight reduction in size towards posterior portion of chin; scales near posterior infralabials flat, subimbricate, and larger than scales in median area of chin; infralabials four, first largest, fourth below center of eye; throat with small granular scales, the posterior region with larger, granular, and pointed scales directed upward (some directed posteriorly); dorsal scales on neck and body small, granular, and juxtaposed; dorsals around midbody and on posterior portion of trunk pointed, granular, or, in frontal view, triangular, mostly directed posteriorly, some scales on flanks and in lumbar region slightly flattened; 40 (36–50) middorsal scales in one HL, 89 between levels of axilla and groin; ventrals flat, smooth, imbricate, with an ovoid posterior margin, increasing moderately in size from gular region to belly, posterior ventral scales longer than wide; 12 (11–15) midventral scales in one HL, 27 between levels of axilla and groin, 32 to border of cloaca; ventral escutcheon patch with 53 (38–46) scales, some of which have slightly pointed posterior margins, 7 (6–7) scales long and 13 (10–13) wide, escutcheon long/wide ratio 54% (50–70%); subfemoral escutcheon scales five (4–5) per thigh, arranged in a single row; transition between ventrals and scales on flanks abrupt; scales around midbody 92 (69–92), of which 18 are ventrals (16–19); scales on precloacal plate similar to ventrals, except for those on border of cloaca, which are smaller; tail dorsally and laterally with flattened, smooth, and imbricate scales, less elongated than ventrals (with a transitional zone at base of tail); underside of tail with a median row of moderately enlarged scales, mostly with a repeated series of one median scale bordered laterodistally by one scale, followed by a slightly larger median scale that is in contact laterodistally with two scales ( Figs. 3 , 7 ), constituting a regular tail sequence of 1'1''; dorsal scales on forelimbs granular; scales on hind limbs flat, smooth, imbricate on anteroventral thigh and shank surfaces, granular elsewhere; fingers, from longest to shortest, IV-III-II-V-I; toes IV- III-II-V-I, fourth and third toes about the same length; lamellae under fourth finger six (5–8), under fourth toe eight (8–9), proximal lamellae slightly larger than distal ones; claws enclosed by an ungual sheath composed of six scales, as typical for the genus.

Hemipenis morphology. The everted hemipenis of SMF 50968 ( Fig. 6 A–B) is a small, bilobate organ, divided for around one third of its length, with a naked base; sulcus spermaticus bordered by well-developed, smooth sulcal lips; ornamentation of papillate calyces present on each lobe, asulcate area of the truncus covered by small spines; a third lobule-like rising from the pedicel, not connected to the sulcus spermaticus, and covered with papillate calyces.

Coloration in preservative (alcohol 70%; variation among the paratypes in parentheses). Dorsal ground color Hair Brown (277); occipital marks Beige (254) (Gray Horn Color (268) in females), posterior margin of orbit bordered with Sepia (279); dorsum of head with small Beige (254) blotches; infra- and supralabials with alternating Sepia (279) and white bars; chin and throat with Sepia (286) marks on a dirty white background; venter slightly pigmented with Vandyke Brown (281); escutcheon scales unpigmented in the center, with Olive Brown (278) borders.

Coloration in life ( Fig. 10 ; variation among the paratypes in parentheses). Dorsal ground color Glaucous (272), with small scattered Lavender Blue (195) and Sepia (286) dots; neck region Olive Brown (278); an indistinct Sepia (286) line from tip of snout to anterior border of eye; two diffuse postorbital Sepia (286) lines, one directed towards the occipital region and the other towards the ear; infra- and supralabials with alternating Sepia (279) and white bars; a vaguely M-shaped dirty white (Smoke Gray (267) in females) occipital mark bordered with Burnt Sienna (38); top of head suffused with Fawn Color (258) and Lavender Blue (195); chin and throat with Sepia (286) reticulations on a Chamois (84) background; venter Lavender Blue (195) suffused with Cinnamon-Rufous (31); an indistinct dorsolateral pale line from behind the ear to mid tail; tail Cinnamon-Rufous (31); a Sepia (286) line from above groin to mid tail.

Distribution and habitat. Lepidoblepharis emberawoundule is currently known from a few sites in eastern Panamanian montane forests and Chocó-Darién moist forests ( Fund 2011), from 227 to 773 m elevation in Darién and Panamá provinces as well as in the Comarcas Emberá and Guna Yala. Most probably, L. emberawoundule lives in the leaf-litter and feeds on small invertebrates like other Lepidoblepharis ( Vitt et al. 2005) .

Etymology. The name emberawoundule is a compound word in honor to “the forest guardians”, the three indigenous peoples inhabiting eastern Panama; embera: Emberá Indians from the foothills of Jingurudó, Bagre, Sapo, Darién, and Pirre mountain ranges; woun: Wounaan Indians, mainly from the Tuira basin and Majé mountain range; dule: meaning people in the language of the Guna Indians from the Caribbean and Pacific versants of the San Blas and Darién mountain ranges.

Figures

FIGURE 3. Morphological distinction among Lepidoblepharis from Lower Central America; images from left to right: dorsals, ventral views of foot, tail, and mental region; scale bars equal 1 mm. (A) L. emberawoundule; (B) L. rufigularis; (C) L. sanctaemartae; (D) L victormartinezi; (E) L. xanthostigma.

FIGURE 4. Feet and hands of Panamanian Lepidoblepharis. Lepidoblepharis emberawoundule: right foot of SMF 81954 and right hand of MHCH 2957; L. rufigularis (SMF 50659): right foot and right hand; L. sanctaemartae: left foot of SMF 97419 and right hand of MHCH 2948; L. victormartinezi: left foot and left hand of SMF 89963; L. xanthostigma: left foot of SMF 90190 and left hand of SMF 89576. Scale bars equal 1 mm, tip of fourth finger and toe marked with an asterisk.

FIGURE 5. Escutcheon scales in males of Panamanian Lepidoblepharis. (A) L. emberawoundule: ventral escutcheon with 46 scales, 7 scales long x 11 wide, and subfemoral escutcheon with 4 scales per side, of SMF 81951 (SVL = 25 mm); (B) L. rufigularis: ventral escutcheon with 62 scales, 7 scales long x 13 wide, and subfemoral escutcheon with 3 and 4 scales per side, of SMF 50659 (SVL = 25 mm); (C) L. sanctaemartae: ventral escutcheon of SMF 97419 (SVL = 22 mm) with 92 scales, 10 scales long x 12 wide; (D) L. victormartinezi: ventral escutcheon of SMF 89963 (SVL = 27 mm) comprising 57 intact scales (an original total of 61 is assumed considering the symmetrical scale arrangement of the non-damaged portions), 7 scales long x 14 wide; (E – F) L. xanthostigma: (E) ventral escutcheon with 75 scales, 12 scales long x 11 wide, and subfemoral escutcheon with 2 scales per side (no additional scales hidden under strings), of SMF 90189 (SVL = 35 mm), and (F) ventral escutcheon with 36 scales, 6 scales long x 8 wide, and no subfemoral escutcheon of AB 1250 (SVL = 24 mm). All scale bars equal 1 mm.

FIGURE 6. Hemipenis morphology of four Lepidoblepharis species from Panama. Sulcate-asulcate view, (A – B) L. emberawoundule SMF 50968; (C – D) L. sanctaemartae SMF 97419 (E – F) L. victormartinezi SMF 50950; (G – H) L. xanthostigma SMF 91558. All scale bars equal 1 mm.

FIGURE 7. Comparison between Lepidoblepharis emberawoundule, L. microlepis (holotype), and L. peraccae (holotype). Mental plate and postmental scales in (A) L. microlepis and (B) L. emerawondule; (C) right foot of L. microlepis; (D) right foot of L. emberawoundule; (E) subcaudal scales in L. microlepis, each of the slightly enlarged median subcaudal scales is bordered laterodistally by only one scale, leading to a regular tail sequence of 1 ' 1 '; (F) subcaudal scales in L. emberawoundule, the larger of the enlarged subcaudals are bordered laterodistally by two scales, the smaller ones by one, forming a regular tail sequence of 1 ' 1 ''; (G) right foot and (H) right hand of the holotype of L. peraccae; scale bars equal 1 mm.

FIGURE 10. Holotype of Lepidoblepharis emberawoundule (male SMF 50968) in life. (A) entire specimen; (B) dorsal view of the head; (C) lateral view of the head.